Monograph of Thysananthus subgenus Thysananthus (Lejeuneaceae, Marchantiophyta)
FURTHER NAMES IN THYSANANTHUS
Thysananthus africanus (Sim) S. Arnell, Bot. Not.: 282. 1953, nom. illeg. [Art. 53.1]
Thysanolejeunea africana Sim, Trans. Roy. Soc. South Africa 15: 50. 1926.—Type:
South Africa. “Perie forest, Kaffraria”, Sim 346 (not seen). Wigginton & Grolle (1996) mentioned: “To be clarified. Collected only once?”.
Thysananthus erosus Steph., Sp. Hepat. 6: 568. 1924.—Type: NEW GUINEA. “Ledermann s.n.” (not seen); Icon. Steph. nr. 10180 = Conoscyphus Mitt.? Verdoorn (1934) considered T. erosus as a member of Jungermannieae. Judging from the original description and the drawing of T. erosus of Stephani’s “Icones Hepaticarum Ineditae”, which shows one leaf and one underleaf, this might be Conoscyphus.
Thysananthus frauenfeldii Reichardt, Verh. K. K. Zool.-Bot. Ges. Wien 16: 958. 1866.
Mastigolejeunea frauenfeldii (Reichardt) Verd., Blumea 1: 230. 1934.—Type: TAHITI. Novara Exped., “…um Papiete, Jelineck s.n.” (not seen).
153 Thysananthus fruticosus fo. pendula Herzog, Trans. Brit. Bryol. Soc. 1: 319.1950, nom. inval.
[Art 32.1(d)]. Material: MALAYSIA. Sarawak: “Dulit, … n. 2277”, 1932, P. W.
Richards s.n. (not seen).
Thysananthus hebridensis Steph., in Stephani & Watts, J. & Proc. Roy. Soc. New South Wales 48: 134. 1914, nom. inval. [Art 32.1(d)]. Material: VANUATU [“New Hebrides”]. Tafea: Aneityum, Gunn s.n. (not seen).
Thysananthus integerrimus Steph., Sp. Hepat. 6: 567. 1924.—Type: NEW GUINEA.
“Ledermann s.n.” (not seen); Icon. Steph. nr. 10187. Verdoorn (1934) considered T.
integerrimus as a member of Jungermannieae.
Thysananthus lacerostipulus Steph., Sp. Hepat. 6: 567. 1924.—Type: NEW GUINEA. Ledermann s.n. (not seen); Icon. Steph. nr. 10223 = Conoscyphus Mitt. ? Verdoorn (1934) considered T. lacerostipulus as a member of Jungermanieae. Judging from the original description and the drawing of T. lacerostipulus of Stephani’s “Icones Hepaticarum Ineditae”, which shows one leaf and one underleaf, this might be Conoscyphus.
Thysananthus polymorphus var. β planifolia (Sande Lac.) Schiffn., Consp. Hep. Archip. Ind.:
305. 1898. Phragmicoma polymorphua fo. β planifolia Sande Lac., Syn. Hepat. Jav.:
58. 1856.—Type: INDONESIA. Java: “ad montem Salak, Zollinger, coll. sub 3560 (pl.
femina); Herb. v. d. Bosch (pl. mascula)” (not seen).
Thysananthus sikkimensis Steph ex Schiffn., Österr. Bot. Z. 49: 205. 1899, nom. inval. [Art 32.1(d)]. Material: BHUTAN. “between Maria Basti & Labar, 5000–6000 ft.”, 12 Apr 1898, R. L. Durel 159 (not seen).
Thysananthus ungulatus (Mitt.) Steph., Sp. Hepat. 6: 567. 1924. Lejeunea ungulata Mitt., J.
Proc. Linn. Soc., Bot. 5: 110. 1861.—Type: BURMA. “in Pegu, ad Rangoon,…”, McClelland s.n. (not seen) = Spruceanthus Verd.? Verdoorn (1934) doubtfully referred T. ungulatus to Spruceanthus (“cf. sub Spruceanthus”). Judging from the original description, this might indeed be Spruceanthus.
154 Thysananthus wardianus (Mitt.) Verd. (“Steph.”), Ann Bryol., Suppl. 4: 166. 1934 as in Index Hepaticarum (Geissler & Bischler, 1990). Lejeunea wardiana Mitt., J. Proc. Linn.
Soc., Bot. 5: 109. 1861.—Type:INDIA.“in Assam superiore”, Griffith s.n. (not seen) = Thysananthus subg. Mastigolejeunea (Spruce) comb. nov., syn. fide Verdoorn (1934): 166.
Thysananthus yokogurensis (Steph.) S. Hatt., J. Hattori Bot. Lab. 8: 32, 1952, nom. inval. [Art 32.1(d)]. Thysanolejeunea yokogurensis Steph. ex Yoshinaga, Bot. Mag. (Tokyo) 15:
93. 1901, nom. inval. [Art 32.1(d)]. Material: JAPAN. (not seen).
ACKNOWLEDGEMENTS
The research was supported by a Royal Thai government scholarship in cooperation with the German Academic Exchange Service (DAAD). This research also received support from the SYNTHESYS Project (http://www.synthesys.info/) financed by the European Community Research Infrastructure Action under the FP7 "Capacities" Program. I am deeply grateful to Prof. S. R. Gradstein for his contribution to the manuscript text and useful comments and suggestions on the manuscript. I would like to thank J. Wang for providing useful information. I express my gratitude to the curators of the following herbaria for making the specimens available and the loan of specimens: BCU, BKF, BM, BM-K, BR, BZ, C, FH, G, GOET, H, HIRO, JE, L, LAE, NICH, NSW, NY, PC, PSU, S, SING, STR, U and W.
Thanks are also due to C. Gehrig-Downie and S. Chantanaorrapint for additional fresh materials, and to B. Raufeisen for the base map of the world.
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159
FIG. 1. Method of measurement in subg. Thysananthus. A. Leaf, ventral view: a = appendage, d = dorsal leaf margin, f = lobule free margin, k = keel, l = leaf length, lb = lobule length, t1,2 = lobule teeth, u = auricle, v = ventral leaf margin, w = leaf width, wb = lobule width. B. Underleaf, ventral view: u = auricle, l = underleaf length, w = underleaf width. C. Perianth, ventral view: b = beak, l = perianth length, w = perianth width. D. Portion of shoot with Lejeunea-type branching, ventral view: c = collar. E. Portion of shoot with androecia, gynoecium and lejeuneoid innovation (dichasial pattern), ventral view.
F. Flagella at stem base, ventral view. G. Lateral merophyte cells arranged in straight longitudinal line: h= hyaline appillae.
H. Underleaf base in longitudinal section. (A from lectotype of T. appendiculatus; B–C from holotype of T. discretus; D from T. spathulistipus, Sukkharak 766 (BKF, GOET); E from T. spathulistipus, Sukkharak 746 (BKF, GOET); F from isolectotype of T. comosus; G from T. spathulistipus, Sukkharak 761 (BKF, GOET); H from holotype of T. discretus).
160
FIG. 2. Leaves and underleaves adnation and appendages in subg. Thysananthus. A. Portions of plant, ventral view: d = adnation, a = appendage. B. Adnation in longitudinal section. C. Appendages at bases of leaf keels. D–F. Female bracts, ventral view: a = appendage. G. Appendage and hyaline papilla at base of bract keels: h = hyaline papilla. H. Appendage at mid-basal portion of bract lobule. I. Perianth, ventral view: a = appendage. J. Appendage and hyaline papilla of perianth: h = hyaline papilla. K. Portions of plant showing appendage at lobule base, ventral view: a = appendage. L. Appendage: b = lobule, a = appendage, h = hyaline papilla, s = stem. (A from isolectotype of T. comosus; B from T. spathulistipus, Sukkharak 766 (BKF, GOET); C, E, H from holotype of T. discretus; D from holotype of T. gottschei var. continuus; F from T.
appendiculatus, Gradstein 3875 (U); G from T. appendiculatus, Streimann & Tamba 12358 (JE); I–J from T. mollis, Gradstein 3795 (U); K from T. retusus subsp. retusus, Sukkharak 733 (BKF, GOET); L from T. montanus, Streimann 13639 (LAE)).
161
FIG. 3. Androecia, Gynoecia and young sporophyte in subg. Thysananthus. A. Portion of androecial spike showing epistatic and hypostatic lobules, ventral view: e = epistatic, h = hypostatic. B. Male bract with 2 antheridia. C. Antheridia. D. Juvenile perianth. E. perianth and protruding archegonial neck. F. Archegium and archgonial neck. G. Later stage of development. H.
Sporophyte prior to elongation of seta. I. Sporophyte after partial elongation of seta. J. Mature perianth, just dehiscence. K.
Transverse section of seta. (A from T. amazonicus, Cornelissen & Steeger C682 (U); B from T. spathulistipus, Gradstein 3964 (U); C from T. spathulistipus, Gradstein 3965 (U); D–G from T. spathulistipus, Sukkharak 766 (BKF, GOET); H–I, K from T. retusus subsp. retusus, Streimann 40547 (JE); J from T. spathulistipus, Sipman 6992 (U)).
162
FIG. 4. Sporophyte in subg. Thysananthus. A. Outer surface of capsule valve, showing rectangular-shaped hinge. B–D. Cells of the outer layer of the capsule valve, surface view. E. Inner surface of dehiscing capsule, showing base of capsule and longitudinal ridges. F. Cells of the inner layer of the capsule valve, surface view. G. Inner surface of dehiscing capsule, showing elaters patterns of either 1+3+5+6 or 2+4+6+6 per valve. H. Elaters. I. Spores. (A–G from T. spathulistipus, Yong &
Ponniah 2981 (GOET); H from T. spathulistipus, Thiers & Roy 2397 (U); I from T. spathulistipus, Sipman 6992 (U)).
163
FIG. 5. Majority rule Bayesian phylogram inferred from the combined chloroplast dataset including differences in sequence lengths (indels). Branch thickness and color indicate bootstrap support and posterior probability values. BPP = posterior probability values from Bayesian analysis, in which gaps treated as missing data; BPPSIC = posterior probability values from Bayesian analysis, in which gaps treated as informative by a simple indel coding strategy; MPB = bootstrap support values from the maximum parsimony optimality criterion, in which gaps treated as missing data; MPBSIC = bootstrap support values from the maximum parsimony optimality criterion, in which gaps treated as informative by a simple indel coding strategy. Modified after Sukkharak et al. (Submitted).
164
FIG. 6.Majority rule Bayesian phylogram inferred from the ITS dataset. For further explanation see Fig. 5.Modified after Sukkharak et al. (Submitted).
165
FIG. 7. Thysananthus aculeatus Herzog. A. Portion of shoot, ventral view. B–C. Leaves, ventral view. D. Underleaves, ventral view. E. Female bracts, ventral view. F. Female bracteole, ventral view. G. Perianth, dorsal view. H. Perianth, ventral view. I. Leaf apices. J. Margin cells of leaf lobe. K. Median cells of leaf lobe. L. Basal cells of leaf lobe. M–N. Leaf lobule apices. O. Cross section of stem. (A–B, D–M, O from the lectotype; C, N from paralectotype).
166
FIG. 8. Thysananthus aculeatus Herzog. A. Leaves, ventral view. B. Perianth, dorsal view. C. Perianth, ventral view. D. Leaf lobule apices. E. Portion of shoot, ventral view. F. Leaves, ventral view. G. Perianth, dorsal view. H. Perianth, ventral view.
I. Leaf lobule apices. (A–D from holotype of T. formosanus; E–I from the holotype of T. richardsianus).
167
FIG. 9. Thysananthus amazonicus (Spruce) Schiffn. A. Portion of shoot, ventral view. B. Leaves, ventral view. C.
Underleaves, ventral view. D. Female bracts, ventral view. E. Female bracteole, ventral view. F. Perianth, dorsal view. G.
Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L.
Leaf lobule apices. M. Cross section of stem. (All from Gradstein s.n. (GOET)).
168
FIG. 10. Thysananthus appendiculatus Steph. A. Portion of shoot, ventral view. B. Leaves, ventral view: a = appendage. C.
Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L. Leaf lobule apices. M. Cross section of stem. N. Portion of shoot, ventral view. O. Leaves, ventral view. P.
Underleaves, ventral view. (A–C, H–M from the lectotype; D–G from Streimann 13859 (JE); N–P from Gradstein 3875 (GEOT)).
169
FIG. 11. Thysananthus combinatus sp. nov. A. Portion of shoot, ventral view. B. Leaves, ventral view. C. Underleaves, ventral view. D. Female bracts, ventral veiw. E. Female bracteole, ventral view. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apex. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L. Leaf lobule apices.
M. Cross section of stem. (All from the holotype).
170
FIG. 12. Thysananthus comosus Lindenb. A. Portion of shoot, ventral view. B–C. Leaves, ventral view. D–E. Underleaves, ventral view. F. Female bracts, ventral view. G. Female bracteole, ventral view. H. Perianth, dorsal view. I. Perianth, ventral view. J. Leaf apex. K. Margin cells of leaf lobe. L. Median cells of leaf lobe. M. Basal cells of leaf lobe. N. Leaf lobule apices. O. Cross section of stem. P. Oil bodies (A, C, E–O from the lectotype, B, D from Schiffner s.n. (GOET), P from Sukkharak 730 (BKF, GOET)).
171
FIG. 13. Thysananthus convolutus Lindenb. A. Portion of shoot, ventral view. B. Leaves, ventral view. C. Underleaves, ventral view. D. Female bracts, ventral view. E. Female bracteole, ventral view. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L. Leaf lobule apices. M. Cross section of stem. N. Oil bodies. (A–M from the lectotype; N from Sukkharak 717 (BKF, GOET)).
172
FIG. 14. Thysananthus convolutus var laceratus (Steph.) comb. nov. A. Portion of shoot, ventral view. B. Leaves, ventral view. C. Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view. F.
Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apex. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K.
Basal cells of leaf lobe. L. Leaf lobule apices. M. Leaf lobule base. N. Cross section of stem. (All from the lectotype).
173
FIG. 15. Thysananthus discretus Sukkharak & Gradst. A–B. Portion of shoot, ventral view. C–D. Leaves, ventral view. E–F.
Underleaves, ventral view. G. Female bracts, ventral view: a appendage. H. Female bracteole, ventral view. I. Perianth, dorsal view. J. Perianth, ventral view. K. Leaf apex. L–M. Margin cells of leaf lobe. N. Median cells of leaf lobe. O. Basal cells of leaf lobe. P. Leaf lobule apices. Q. Cross section of stem. (A, C, E, G–K, M–Q from the holotype; B, D, F, L from Streimann 13622 (JE)).
174
FIG. 16. Thysananthus gottschei (Jack & Steph.) Steph. A. Portion of shoot, ventral view. B. Leaves, ventral view. C.
Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apex. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L. Leaf lobule apex. M. Cross section of stem. (A–C, H–M from the lectotype; D–G from Gradstein 10324 (GOET)).
175
FIG. 17. Thysananthus gottschei var. continuus var. nov. A. Portion of shoot, ventral view. B. Leaves, ventral view: a = appendage. C. Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view.
F. Perianth, dorsal view. G. Perianth, ventral view. H. Margin cells of leaf lobe. I. Leaf lobule apices. J. Leaf lobule base.
(All from the lectotype).
176
FIG. 18. Thysananthus spathulistipus (Reinw. et al.) Lindenb. A. Portion of shoot, ventral view. B. Leaves, ventral view. C.
Underleaves, ventral view. D. Female bracts, ventral view. E. Female bracteole, ventral view. F. Perianth, dorsal view. G.
Perianth, ventral view. H. Leaf apex. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L.
Leaf lobule apices. M. Cross section of stem. N. Oil bodies. (A–M from the holotype; N from Sukkharak 716 (BKF, GOET)).
177
FIG. 19. Thysananthus spathulistipus (Reinw. et al.) Lindenb. A–D. Portion of shoot, ventral view. E–H. Leaves, ventral view. (A, E from the holotype of T. spathulistipus var. borneensis; B, F from holotype of T. minor; C, G from holotype of T.
fuscobrunneus; D, H from holotype of T. triquetrus).
178
FIG. 20. Thysananthus anguiformis (Hook. f. & Taylor) Steph. A. Portion of shoot, ventral view. B. Leaves, ventral view: a
= appendage. C. Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L. Leaf lobule apices: h = hyaline papilla. M. Cross section of stem. (All from the lectotype).
179
FIG. 21. Thysananthus pancheri (Steph.) Hürl. A. Portion of shoot, ventral view. B. Leaves, ventral view. C. Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view. F. Perianth, dorsal view. G.
Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L.
Leaf lobule apices: h = hyaline papilla. M. Cross section of stem. (All from the lectotype).
180
FIG. 22. Thysananthus fruticosus (Lindenb. & Gottsche) Schiffn. A. Portion of shoot, ventral view. B. Leaves, ventral view:
a = appendage. C. Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apex. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe, showing the vitta. L. Leaf lobule apices. M. Cross section of stem. (A–C, H–M from the lectotype; D–G from Brass 22185 (JE)).
181
FIG. 23. Thysananthus fruticosus (Lindenb. & Gottsche) Schiffn. A–B. Portion of shoot, ventral view: a = appendage. C–D.
Leaves, ventral view: a = appendage. E. Underleaves, ventral view. (A, C, E from Beccari s.n. (JE); D, E from Streimann 25733 (JE)).
182
FIG. 24. Thysananthus mollis Steph. A. Portion of shoot, ventral view. B. Leaves, ventral view. C. Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view. F. Perianth, dorsal view: a = appendage. G. Perianth, ventral view, a = appendage. H. Leaf apex. I. Margin cells of leaf lobe. J. Median cells of leaf lobe.
K. Basal cells of leaf lobe. L. Leaf lobule apices. M. Cross section of stem. (All the holotype).
183
FIG. 25. Thysananthus mollis Steph. A–C. Portion of shoot, ventral view. D–F. Leaves, ventral view. G–I. Underleaves, ventral view. (A, D, G from Streimann & Bellamy 13049 (JE); B, E, H from Streimann 26631 (JE); C, F, I from Gradstein 3792 (GOET)).
184
FIG. 26. Thysananthus montanus Gradst. et al. A. Portion of shoot, ventral view. B. Leaves, ventral view. C. Underleaves, ventral view. D. Female bracts, ventral view. E. Female bracteole. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L. Leaf lobule apex. M. Cross section of stem. (All from the holotype).
185
FIG. 27. Thysananthus retusus (Reinw. et al.) B. Thiers & Gradst. A. Portion of shoot, ventral view. B. Leaves, ventral view.
C. Underleaves, ventral view. D. Female bracts, ventral view. E. Female bracteole, ventral view. F. Perianth, dorsal view. G.
Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe, showing the vitta. L. Leaf lobule apices. M. Cross section of stem. N. Oil bodies, non-vittate cells. O. Oil bodies, vittate cells.
(A–M from the holotype; N–O from Sukkharak 733 (BKF, GOET)).
186
FIG. 28. Thysananthus retusus (Reinw. et al.) B. Thiers & Gradst. A–E. Portion of shoot, ventral view. F–I. Leaves, ventral view. J–L. Underleaves, ventral view. M–N. Leaf lobule apices. (A, F, J, M from the holotype of T. australis; C, H, K, N from Gradstein 3963 (GOET); D, from the holotype; E, I, L from Degelius 223 (JE).—T. retusus subsp. sellingii (Hürl.) comb. nov. B. Portion of shoot, ventral view. G. Leaves, ventral view. (All from the holotype).
187
FIG. 29. Distribution of subg. Thysananthus species. A. Thysananthus amazonicus (Spruce) Schiffn. B. Thysananthus spathulistipus (Reinw. et al.) Lindenb. C. Thysananthus mollis Steph., Thysananthus montanus Gradst. et al. D.
Thysananthus pancheri (Steph.) Hürl. E. Thysananthus anguiformis (Hook. f. & Taylor) Steph. F. Thysananthus comosus Lindenb. G. Thysananthus fruticosus (Lindenb. & Gottsche) Schiffn. H. Thysananthus aculeatus Herzog. I. Thysananthus appendiculatus Steph. J. Thysananthus retusus (Reinw. et al.) B. Thiers & Gradst. K. Thysananthus convolutus Lindenb., Thysananthus gottschei (Jack & Steph.) Steph. L. Thysananthus combinatus sp. nov. M. Thysananthus discretus Sukkharak
& Gradst
188 DECLARATION OF THE AUTHOR’S OWN CONTRIBUTION TO THE PAPERS
I am the overall author of all manuscript presented in this thesis. I did not use any other sources as the ones indicated in the chapters. This thesis has not been submitted previously in any form for another degree at any university or institution.
Phiangphak Sukkharak Göttingen, 27 April 2011
189 CURRICULUM VITAE
Name Phiangphak Sukkharak
Born 7 March 1982, Nan, Thailand
05/1998 – 03/2001 Yupparaj Wittayalai School, Chiang Mai, Thailand supported by the Development and Promotion of Science and Technology talents project (DPST), a Royal Thai government scholarship.
05/2001 – 03/2005 B.Sc. in Biology, Chiang Mai University, Chiang Mai, Thailand. (2nd Class Honors) supported by the Development and Promotion of Science and Technology talents project (DPST), a Royal Thai government scholarship.
05/2005 – 03/2008 M.Sc. in Botany, Chulalongkorn University, Bangkok, Thailand supported by the Development and Promotion of Science and Technology talents project (DPST), a Royal Thai government scholarship.
08/2008 – 06/2011 Ph.D in Biodiversity and Ecology, University of Göttingen supported by a Royal Thai government scholarship in cooperation with the German Academic Exchange Service (DAAD).
07/2011 onwards Lecturer at Faculty of Science, Burapha University, Chonburi,
07/2011 onwards Lecturer at Faculty of Science, Burapha University, Chonburi,